Jaguar
Panthera onca (Linnaeus, 1758)
IUCN RED LIST (2008): Near Threatened
Head-body length: ♀ 116−219cm, ♂ 110.5−270cm
Tail: 44−80cm
Weight: ♀ 36.0−100.0kg, ♂ 36.0−158.0kg
Taxonomy and phylogeny
The Jaguar is one of the ‘big cats’ in the genus Panthera and is thought to be most closely related to the Leopard and the Lion with a common ancestor estimated at approximately 3–3.5 million years ago. Historically, eight Jaguar subspecies were recognised, based chiefly on superficial variation in the skull, but later analysis concluded this was insufficient to distinguish populations. Similarly, genetic analysis has shown relatively little differentiation among Jaguars and instead suggests there are gradual genetic changes across a cline from north to south without strong boundaries between populations. Not surprisingly, the greatest differences are found between Jaguar populations at the latitudinal extremes of their range. The same genetic analysis shows four weakly separated regional groups that display some genetic partitioning but not to the extent normally considered to distinguish subspecies. The main division is along the Amazon River, with the largest group comprising all Jaguar populations south of the Amazon. Populations north of the Amazon are divided into three weakly defined groups, northern South America, southern Central America and Guatemala-Mexico. It is thought that the Jaguar underwent a recent and rapid population expansion across the Americas approximately 300,000 years ago, after which there were few barriers impeding constant exchange of genetic material across the range. Even the Amazon and the Andes have apparently been insufficient to completely isolate Jaguar populations. Importantly, both morphological and genetic analyses are based on very small samples across a massive, continental range, and somewhat greater differentiation between populations may be uncovered with more detailed analysis.
Description
The Jaguar is the world’s third largest cat and is the most robustly built of all living cats, analogous only to extinct sabre-toothed felids such as Smilodon. The thickset body has very muscular forequarters, a very deep chest and foreshortened waist. The limbs are short and stout, such that shoulder height is comparable to or only slightly greater than the much more lightly built Leopard. The feet are very broad and rounded with distinctly stubby and splayed digits, especially in the forefeet that act effectively to spread weight on sodden ground and as swimming paddles. The tail is relatively short compared to other large cats, around half the head-body length. The head is short, rounded and massively built, such that males especially have an unusually heavy-headed ‘Pit Bull’ appearance. The Jaguar’s body size is broadly correlated with changes in latitude across a north–south gradient. The smallest Jaguars occur in Mesoamerican forest, with a weight range of 36−51kg (females) and 48−66kg (males) for animals in Mexico and Belize. The largest Jaguars inhabit the wet, savanna woodlands of Brazil (Pantanal) and Venezuela (Los Llanos) with a weight range of 51−100kg (females) and 68−158kg (males). Animals in similar habitat in this region (for example, from the Bolivian and Paraguayan Pantanal) are likely to be similarly large but measurements are mostly lacking.
The Jaguar’s background colour varies in shades of buff-grey, yellow, cinnamon and tawny-orange with crisp white or creamy-white underparts. The body is covered with large, black block-like markings or rosettes with a darker brown interior and usually small black spots inside; small black spots are sometimes also interspersed between the large markings. The lower limbs and underparts are covered in large, solid black spots and blotches while smaller solid spots cover the shoulders, head and face. The ears are short and rounded with black backs and an off-white central patch. Melanism occurs as a dominant inherited trait, with the same pattern of rosettes apparent in oblique light. Melanistic individuals are most common in lowland tropical forest and are less common north of the Amazon River. The incidence of melanism appears to decline towards the periphery of the range, such that it is not known from the Atlantic Forest biome in northern Argentina and south-western Brazil.
Similar species The Leopard has a close resemblance, though the two species are not sympatric in the wild. The Leopard is less heavily built with a less massive, blocky head, and has smaller, rosette-type markings typically lacking the distinctive interior spots found in the Jaguar. The unicolour Puma is the only sympatric large cat and is never melanistic; the only large black Latin American cat is the Jaguar.
The Jaguar is the most aquatic of all the big cats and inhabits some areas which are seasonally inundated for months on end such as the Brazilian Pantanal (shown here) and the flooded forest habitat known as várzea in the Amazon.
Distribution and habitat
The Jaguar occurs from northern Mexico to northern Argentina. It is widely and mostly continuously distributed in northern and central South America east of the Andes, from the northern Andes in Colombia to the Brazilian highlands; it is patchily distributed at the eastern and southern extent of the range in northern Argentina, south-eastern Brazil and Paraguay. Its range in Central America is fragmented with tenuous links between large forested tracts that largely follow the mountainous cordilleras and associated lowlands along the Caribbean coast. The range is largely contiguous in southern Mexico and bifurcates going north, following the Sierra Madre Oriental and Sierra Madre Occidental mountain ranges. Resident, breeding populations no longer occur in the US, but individuals intermittently appear in border areas of Arizona and New Mexico from the northernmost breeding population 150km south of the border in Sonora, Mexico; three individuals, all of them male, have been documented in the US since 2001. The last record of a wild female Jaguar in the US is from 1963. The Jaguar is extinct in El Salvador and Uruguay.
The Jaguar occurs in a variety of forested and wooded habitats. It reaches highest densities in dense subtropical and tropical lowland forest, and in seasonally flooded savanna woodlands (for example, the Pantanal), but it also occurs widely in dry forest, moist and arid areas, dense scrubland, wooded grasslands and mangrove swamps. Jaguars are closely associated with water and inhabit sodden and seasonally inundated habitats; they are excellent swimmers that routinely traverse rivers including greater than 2km widths of the Amazon and Japurá Rivers. They easily cross the Panama Canal. In arid parts of the range (for example, northern Mexico–southern US and eastern Brazil), Jaguars occur in vegetated scrubland and dry woodland associated with watercourses and mountainous terrain. Jaguars shun open landscapes with poor cover including most grassland habitats, though they occur in forested patches and riverine areas throughout some grasslands. Jaguars occur from sea level typically to 2,500m, and are rarely found in montane forest at higher elevations. They are not found above 2,700m in the Andes nor do they occur on central Mexico’s high plateau. With the exception of extensive ranching areas with habitat and wild prey (such as the Pantanal), Jaguars do not occupy heavily modified, anthropogenic habitats. They have been recorded in pine and oil-palm plantations that retain forested fragments or are close to forested habitat.
A rare photograph of a wild melanistic Jaguar, taken in the Amazon forest, Yasuni National Park, Ecuador. Melanism occurs in at least 14 species of wild cats, the adaptive significance of which is still unknown. The trait may be adaptively neutral in which it carries no significant advantage or disadvantage.
A female Jaguar with her five-month-old cub. Compared to male Jaguars, females seem less tolerant of human-modified habitats. Radio-collared females in Mexico’s Calakmul Biosphere Reserve prefer habitat with fewer roads and that is less impacted by cattle ranching and agriculture than habitat used by males.
Feeding ecology
Jaguars have a diverse diet with at least 86 recorded prey species. Like all large cats, they focus on common, large-bodied prey and they are able to kill the largest available mammals, both native (Marsh Deer and tapirs) and introduced (cattle). However, the natural absence of high densities of large deer, antelope or wild cattle species in Latin America means that Jaguar diet typically includes greater proportions of both smaller species and of non-mammals than in other large cats. Across the range, Capybaras, Collared Peccaries and White-lipped Peccaries are usually the most important wild prey species where they occur in sufficient numbers; the distribution of these three species collectively overlaps that of the Jaguar almost exactly. The northern periphery of Jaguar range is the only area where abundant White-tailed Deer dominate the diet, for example 54 per cent of prey by biomass in the Chamela-Cuixmala Biosphere Reserve, south-western Mexico. Reptiles form a bigger part of Jaguar diet than that of any other large cat. In regions where medium-large reptiles are abundant and/or where Capybaras and peccaries are less available, reptiles may be the most important prey, for example Yacaré, Spectacled and Black Caimans in Amazon forest and some parts of the Pantanal; Jaguars take all life-stages of these three species, including large adults and eggs. Jaguars also prey frequently on at least another 14 reptile species, mostly large and medium-sized species. Large freshwater turtles (genus Podocnemis) and terrestrial tortoises (genus Chelonoidis) are important prey in Amazonian flooded forest and Brazilian Atlantic Forest (recorded in up to 25 and 20 per cent of scats, respectively). Jaguars are known to kill four species of marine turtles: Green, Leatherback, Hawksbill and Olive Ridley, chiefly when females are nesting in large aggregations on beaches. Green Turtles appear to be the most important prey species during the nesting season (June–October) in Tortuguero National Park, Costa Rica; a minimum 672 adult Green Turtles (plus one Leatherback and three Hawksbills) were killed by Jaguars in 2005–2010. Snakes, including large anacondas and Boa Constrictors, are also killed by Jaguars although they do not contribute significantly to intake.
Depending on the relative availability of these two primary prey categories – medium-sized mammals and medium to large reptiles – Jaguars also eat a variety of smaller vertebrates: mammals such as primates, armadillos, tamanduas, agoutis and marsupials, and reptiles, chiefly iguanas and tegu lizards. During a period in which peccaries were rare in the Cockscomb Basin, Belize, Jaguars ate mainly Nine-banded Armadillos, Spotted Pacas and Red Brocket Deer. Following 20 years of protection at this site, armadillos are still the most important prey species (42 per cent of biomass consumed) followed by peccaries (15.6 per cent), which are now more common. Similarly, in the Maya Biosphere Reserve, Guatemala, Nine-banded Armadillos and White-nosed Coatis are the most important prey species – 58.1 per cent of biomass for areas hunted by people and 46.3 per cent in better protected areas (where peccaries are more common); peccaries comprise 15.5 per cent of biomass and 27.2 per cent respectively. In Amazonian flooded forest, Jaguars feed mainly on caimans, Brown-throated Sloths and primates. The Giant Anteater is the most frequent prey item in cerrado savanna, central Brazil (Emas National Park) and in dry caatinga thorn-scrub of north-eastern Brazil. As the largest mammalian predator in Latin America, Jaguars kill a wide variety of other carnivores including records of Puma, Ocelot, Margay, Maned Wolf, Crab-eating Fox, Grey Fox, Tayra, skunks, Kinkajou, Cacomistle, olingos, raccoons and coatis. Killed carnivores are frequently abandoned without eating and rarely feature as important prey with the exception of procyonids; that is, White-nosed Coati, South American Coati and Crab-eating Raccoon, which comprise 5–21.5 per cent of biomass consumed in Mesoamerica and some areas in Brazilian Atlantic Forest and the Pantanal. Cannibalism occurs rarely including cases of cubs consumed by infanticidal males, and one incident in which two adult males killed and partially ate an adult female (southern Pantanal, Brazil). Jaguars incidentally eat birds, amphibians and fish. Rheas are important prey (13 per cent occurrence in scats) for Jaguars in the cerrado savannas, central Brazil. Jaguars in the Amazon are recorded killing freshwater dolphins as they fish in shallow water with records from Brazil and Colombia. Jaguars readily kill domestic livestock, chiefly cattle and occasionally horses. In ranching habitats such as the Pantanal and Los Llanos, cattle form a major prey species, and are the most important prey for Jaguars in the southern Pantanal, Brazil (31.7 per cent of kills), and in north-eastern Sonora, Mexico (57.7 per cent of biomass consumed). Jaguars infrequently take domestic pigs from villages, and free-ranging dogs are killed, for example in forest on the Yucatán Peninsula near human communities. Jaguars almost never hunt people; most recorded attacks are caused by extreme provocation, such as during Jaguar hunts, and verified unprovoked attacks are extremely unusual.
Jaguars forage chiefly at dusk, night and early morning. Jaguars hunt primarily on the ground – they are poorly adapted to pursue prey arboreally – as well as in and around water sources. The Jaguar is perhaps the most water-adapted felid and actively hunts in water. Jaguars pursue fleeing prey into water and launch spectacular leaping attacks from high riverbanks onto caimans and Capybaras in the water below. They also search for prey by passively floating downstream with the current to stealthily locate caimans and Capybaras resting on river beaches, and they readily launch attacks directly from the water. The Jaguar has a massively constructed skull equipping it with proportionally the strongest bite force of all the large cats. Although prey may be killed with a typically feline suffocating throat bite, Jaguars are unusual among cats for often killing large prey (including very large caimans and cattle) by a crushing bite to the skull, typically delivered at the rear of the braincase. This enormously powerful bite also allows Jaguars to open the carapace of large freshwater turtles and land tortoises. They kill large marine turtles by biting the vulnerable neck close to the skull.
Hunting success of Jaguars is unknown. They drag kills into dense cover, with anecdotal reports that dragging is more frequent and occurs over large distances in ranching landscapes where Jaguars are persecuted. A small female Jaguar with an estimated weight of 41kg dragged a 180kg heifer 200m through a densely forested ravine in Venezuela. Jaguars are not recorded covering kills. They return to large kills, which are consumed over a number of days. Jaguars readily scavenge, particularly from dead livestock that are the chief source of carrion in much of the range. Two different male Jaguars were recorded feeding on the carcass of a beached marine dolphin (species uncertain) on the northern Honduras coast.
A Jaguar narrowly misses a Capybara during a daylight hunt in the Brazilian Pantanal. In habitats with marked seasonality, Jaguars focus their hunting efforts along river systems during the dry season.
Unusual among felids, the Jaguar often kills large prey with a crushing bite to the skull or nape. Recent film of a Jaguar killing a large caiman using this technique shows the reptile paralysed instantly by the bite – a safe and efficient method for dealing with powerful, dangerous prey.
Social and spatial behaviour
The socio-spatial ecology of the Jaguar is still relatively poorly known, though they clearly follow the fundamental felid pattern of being largely solitary and territorial. Both sexes maintain enduring territories in which the ranges of males are larger than those of females. Exclusive range use appears to be limited to small core areas; adults in some populations have highly overlapping ranges, possibly due to marked seasonal changes in the distribution of water and hence prey. Pantanal females establish largely exclusive ranges during the wet season, but they overlap significantly in the dry season while males overlap extensively in both seasons. Similarly, Cockscomb Basin (Belize) males have highly overlapping ranges with some camera-trap locations visited by as many as five males per month.
As is typical for felids, male ranges overlap numerous female ranges, but high overlap among males means the reverse is also true for some populations. In one southern Pantanal study, one adult female’s range was overlapped by at least three adult males, and her range was entirely encompassed by one of them. The following year, she overlapped with two males (one of them from the previous year and one new male); these same two males also overlapped a second adult female’s range extensively.
Even with greatly overlapping ranges, it seems that adult Jaguars are no more sociable than other largely solitary cats. Adults engage in typically territorial behaviours, such as roaring and urine-marking, perhaps serving more to avoid encounters rather than to demarcate exclusivity. Of 11,787 telemetry locations recorded over 3.5 years during the southern Pantanal research, there were 32 possible interactions between a female and a male jaguar, 21 possible encounters between two males, and only one possible encounter between two females. Two adult males that were clearly not siblings shared a feral pig carcass. Males are sometimes seen interacting amicably with females and their cubs, presumably when the male is the putative sire. Aggressive interactions between adults appear to be rare. Of 697 camera-trap images of 23 individual male jaguars in Cockscomb Basin, Belize, only three photographs showed serious flesh wounds or scars that could have been inflicted by another male. Nonetheless, there are occasional records of fatal fights; for example, an adult male was killed by another male presumably in a clash over territory in the southern Pantanal, Brazil.
Territory size varies with habitat quality and prey availability, from 30−47km2 (females in moist savannas, Venezuela and Brazil) to 1291km2 (male, dry savanna, Paraguay). Range size estimates are confounded by small sample sizes and few studies using GPS telemetry (which avoids the problem of underestimating range size that is common in conventional telemetry studies). Of studies with large samples and/or from GPS telemetry, average range sizes are available from wet Pantanal savanna woodland (females 57−69km2; males 140−170km2), Atlantic Forest, Brazil (females 92−212km2; males 280−299km2) and dry Gran Chaco savanna woodland, Paraguay (females 440km2; males 692km2). A lone male in arid lowland desert and pine-oak woodlands in Arizona used at least 1,359km2 between 2004 and 2007 (calculated by camera-trapping and hence an underestimate). Two males in tropical forest in the southern Yucatán Peninsula, Mexico, had ranges of at least 1,000km2. Home range sizes in seasonally inundated habitat (for example, the Pantanal) often contract in the wet season when flooded areas limit the space available to prey. The density of Jaguar populations increases along a cline loosely associated with precipitation. The lowest density populations are found in the arid regions of the range such as in Sonora, Mexico (1.05 Jaguars per 100km2, Northern Jaguar Reserve, Mexico) and caatinga habitat, north-eastern Brazil (2.7 Jaguars per 100km2, Parque Serra da Capivara, Brazil). The highest densities are recorded from very wet lowland forest (7.5−8.8 Jaguars per 100km2, Chiquibul and Cockscomb, Belize) and wet savanna woodlands (6−7 Jaguars per 100km2 and possibly as high as 11 Jaguars per 100km2, Pantanal, Brazil).
A male Jaguar and the adult female Leatherback Turtle it has killed, with Black Vultures in attendance, Tortuguero National Park, Costa Rica. Despite their size, nesting marine turtles are powerless to defend themselves against Jaguars.
A female Jaguar and her two-month-old cubs cool off in Rio Cuiabá in the Brazilian Pantanal at the height of the dry season. There is almost no information on the survival and dispersal of wild Jaguar cubs.
Reproduction and demography
Reproductive patterns from the wild are very poorly known. Although it is common to find published references to a mating season in the Jaguar, reproduction is aseasonal. It is possible there are birth peaks in regions with marked seasonal differences, for example in Los Llanos and the Pantanal which have marked wet and dry seasons. Oestrus lasts 6−17 days and gestation lasts 91−111 days, averaging 101−105 days. Litter size is one to four cubs, averaging two (captivity). Weaning begins around 10 weeks and suckling typically ceases by four to five months. Cubs reach independence at 16−24 months. Dispersal is poorly known but it appears to be typically feline in which females settle close to their natal range while males disperse more widely. Inter-litter interval is unknown from the wild. Both sexes are sexually mature at 24−30 months; females first give birth at 3−3.5 years and can reproduce to 15 years (captivity). It is not known when wild males first breed but as for other large cats, it is likely to be no earlier than the age at which males are first able to become territorial; that is, no younger than three to four years old.
Mortality Mortality rates and factors in wild Jaguars are largely unknown. Adult jaguars have no natural predators and are killed principally by people and rarely by other Jaguars in territorial disputes. Adult Jaguars on the boundary of Calakmul Biosphere Reserve, Mexico, were close to human communities and were exposed to domestic carnivore diseases (feline heartworm and toxoplasmosis) while Jaguars deep inside the reserve were not; it is unknown if this led to mortalities. Predators of cubs are poorly known. Infanticide by male Jaguars is recorded, and there is one record of a female killing an unrelated cub at a cow carcass being used by the cub’s mother and the infanticidal female in the Brazilian Pantanal.
Lifespan Poorly known from the wild, but is unlikely to exceed 15−16 years, and reaches 22 years in captivity.
STATUS AND THREATS
Jaguars have been extirpated from an estimated 49 per cent of their historical range and are extinct in El Salvador, the US and Uruguay. Despite this, the Jaguar’s remaining range contains large areas of essentially continuous habitat, in part because the massive forested basins of South America have remained mostly inaccessible until recently. The most extensive Jaguar stronghold with a high probability of long-term persistence is the Amazon basin rainforest, and neighbouring areas of the Pantanal and Gran Chaco woodlands. Large fragments of tropical lowland forest in Central America are also considered to be strongholds, chiefly the Selva Maya of Mexico, Guatemala and Belize; the Río Plátano forest on the border of Honduras and Nicaragua; and a narrow strip of the Chocó-Darién moist forest from northern Honduras through Panama to northern Colombia. Most of the remaining range in Central America and Mexico is highly threatened. Similarly, populations at the periphery of remaining Jaguar range are regarded as severely threatened, particularly those in coastal dry forest in Venezuela; the Gran Sabana woodlands of Guyana, Venezuela and northern Brazil; the Atlantic Forest and cerrado of Brazil; and the Gran Chaco woodlands in northern Argentina. Populations in the inter-Andean valleys in Colombia represent the critical connection between Central America and South America, and are severely threatened.
Latin America has very high rates of habitat conversion for forestry, livestock and agriculture that directly threaten the Jaguar. This is combined with intense persecution from ranchers and pastoralists in livestock areas despite the fact that many cattle losses blamed on predation occur from other factors. People remove very large numbers of Jaguars in some ranching landscapes; for example, an estimated 185−240 large cats (Jaguars and Pumas) were killed between 2002 and 2004 in a 34,200km2 ranching landscape in Alta Floresta, Brazil. Human hunting of prey is also likely to impact Jaguar populations and may be underestimated in areas of intact forest with human communities who rely heavily on the same prey species as large cats. There is also emerging evidence that the species is hunted to supply the Chinese medicinal trade, though this is poorly quantified. Sport-hunting is illegal in all range states, though illegal hunting is popular as recreation in some areas, for example the Pantanal and Los Llanos. There is no commercial hunting of Jaguars for their pelts since international markets were closed in the mid-1970s, though there is still widespread local demand for Jaguar paws, teeth and skins. CITES Appendix I. Red List: Near Threatened. Population trend: Decreasing.